Enzyme variation at morphological boundaries in Maniola and related genera (Lepidoptera: Nymphalidae: Satyrinae)

Abstract

The evolutionary biology of 14 species of Maniolini (Nymphalidae: Satyrinae) was studied. Electrophoretic analysis of 35 enzyme loci identified a larger number of alleles than an1 levels of polymorphism similar to those found in other Lepidoptera. In Maniola jurtina, some populations exhibited a massive heterozygote deficit and sex associated allele frequency differentiation at the GOT-l locus. Allele frequencies in pre- and post-aestivation jurtina from southern Europe were significantly different. At some loci, significant annual differences in allele frequencies were noted. A significant correlation between geographic and genetic distance in allele frequencies was observed, but no correlation was detected between heterozygosity and land area in insular populations. Cluster analysis and nonmetric multidimensional scaling per~ormed on electrophoretic data from populations of Maniola jurtina revealed a dichotomy between 'Eastern' and 'Western' subspecies groups. The analysis of Manioline species fitted existing taxonomies. Genetic differences between Maniola species were much smaller than those between Pyronia and Hyponephele species. Ultrastructural studies of the Maniola Jullien organs revealed a species-specific tooth pattern on the inner surfaces. It is suggested that these structures may be sound production mechanisms of great evolutionary significance to the species. Maniolini ova were studied and it is suggested that their form and chorionic sculpturing owe much to selection induced by oviposition strategy. Chaetotaxy of first instar larvae was undertaken and morphometric analysis of setal lengths was found to be useful, but not unambiguous. Multivariate analysis of chaetotaxy data showed a significant correlation with electrophoretic data. viii The evolution and zoogeography of Maniola is discussed. It is suggested that disjunction, founder effect, rapid post-glacial colonisation and bottlenecking have played a major roles in effecting rapid speciation. It is further suggested that all Maniola species are very recent, perhaps having evolved within the last 50,000 years, and some species almost certainly have evolved in postglacial times •